We have examined the developmental properties of the polydactylous chicken mutant, talpid(2). Ptc, Gli1, Bmp2, Hoxd13, and Fgf4 are expressed throughout the anteroposterior axis of the mutant limb bud, despite normal Shh expression. The expression of Gli3, Ihh, and Dhh appears to be normal, suggesting that the Shh signaling pathway is constitutively active in talpid(2) mutants. We show that preaxial talpid(2) limb bud mesoderm has polarizing activity in the absence of detectable Shh mRNA. When the postaxial talpid(2) limb bud (including all Shh-expressing cells) is removed, the preaxial cells reform a normal-shaped talpid(2) limb bud (regulate). However, a Shh-expressing region (zone of polarizing activity) does not reform; nevertheless Fgf4 expression in the apical ectodermal ridge is maintained. Such reformed talpid(2) limb buds develop complete talpid(2) limbs. After similar treatment, normal limb buds downregulate Fgf4, the preaxial cells do not regulate, and a truncated anteroposterior deficient limb forms. In talpid(2) limbs, distal outgrowth is independent of Shh and correlates with Fgf4, but not Fgf8, expression by the apical ectodermal ridge. We propose a model for talpid(2) in which leaky activation of the Shh signaling pathway occurs in the absence of Shh ligand.
The relationship between physical ability and physical activity is reinforced with this study. The impact of stroke on the family, particularly on time demands of the primary caregiver, suggests the needs of the care partner may not be adequately addressed in the rehabilitation process.
The talpid2(ta2) chick mutant is of interest for the study of limb pattern formation. Talpid2 is a simple Mendelian recessive lethal mutation which affects the mesoderm and results in short, spade-like, polydactylous wings and legs. Here, we describe ta2 limb development with particular attention to those aspects of ta2 which may illuminate the process of normal limb development. From the onset of budding, ta2 limb buds are significantly wider than normal buds along the anteroposterior axis. They lack the normal anterior and posterior necrotic zones and have variable development of the central opaque patch. Interdigital programmed cell death is variable and may result in development of distal phalanges without more proximal ones. Talpid2 wing vasculature is similar to that of normal wings; but ta2 legs are supplied by four large blood vessels. Feathers form regular, parallel rows, similar to normal feathers, but ta2 embryos lack the loose mesenchyme which separates the feather buds. Finally, and most significantly, ta2 wings and legs display anteroposterior polarity. Anterior and posterior limb skeletal elements can be clearly distinguished from one another within the ta2 phenotype. Our observations suggest that the ta2 mutant may be useful in analyzing programmed embryonic cell death and anteroposterior limb pattern formation.
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